CUCURBITALES Dumortier Main Tree, Synapomorphies

Cucurbitopsida Brongn., Enum. Plant. Mus. Paris: xxx, 116. 12 Aug 1843 [’Cucurbitineae’]; Cucurbitanae Reveal in Phytologia 76: 4. 2 Mai 1994

Habit Monoecious, andromonoecious, gynomonoecious, polygamomonoecious, dioecious, androdioecious, or gynodioecious (rarely bisexual), evergreen or deciduous trees, shrubs, lianas, suffrutices, or perennial (rarely annual) herbs.

Vegetative anatomy Root nodules containing nitrogen-fixing endosymbiotic actinobacteria (Frankia) often present; Frankia infection sometimes via intercellular penetration. Phellogen ab initio superficially or deeply seated. Secondary lateral growth normal or anomalous (from concentric cambia or simple cambial cylinder). Cambial initials storied, fusiform. Vessel elements with usually simple (sometimes scalariform) perforation plates; lateral pits alternate or scalariform, simple or reduced bordered pits. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays multiseriate, homocellular or heterocellular, or absent. Axial parenchyma apotracheal, diffuse or diffuse-in-aggregates, or paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent or vasicentric, or banded. Intraxylary phloem usually present. Sieve tube plastids usually Ss type (sometimes S₀ type). Nodes 1:1 or 1:3, unilacunar with one or three leaf traces, 3:1–3, trilacunar with one to three traces, or 5:5, pentalacunar with five traces. Heartwood sometimes with resinous substances. Branched idioblasts with bitter substances. Calciumoxalate as prismatic crystals, druses and raphides.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple or branched, sometimes bristle-like, candelabra-shaped, stellate or prickles; glandular hairs often present, sometimes lepidote glands and pearl glands; cell walls of hairs often calcified; calcified cystoliths and similar structures often present at hair bases or in adjacent cells.

Leaves Usually alternate (spiral or distichous, sometimes opposite, rarely verticillate), simple or usually palmately (rarely pinnately) compound, entire or palmately (rarely pinnately) lobed, with conduplicate ptyxis. Stipules present (sometimes large, cauline-extrapetiolar or intrapetiolar) or absent; leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Venation pinnate or palmate. Stomata anomocytic, helicocytic, anisocytic, or paracytic. Cuticular wax crystalloids usually absent (sometimes as rosettes of platelets, Fabales type). Mesophyll often with sclerenchymatous idioblasts. Cystoliths usually present. Leaf margin entire, crenate, dentate or serrate; leaf teeth often begonioid or cucurbitoid.

Inflorescence Axillary, panicle, thyrse, raceme, spike, bostrycoid, dichasial, or flowers solitary. Floral prophylls (bracteoles) often absent.

Flowers Usually actinomorphic (sometimes zygomorphic or bisymmetrical). Hypanthium often present. Usually epigyny (sometimes hypogyny or half epigyny). Sepals two to five (to 16), with imbricate, valvate or open aestivation, free from each other or more or less connate (rarely absent). Petals two to five (to 16), with valvate, induplicate-valvate or imbricate (sometimes open) aestivation, in lower part often connate into campanulate corolla (rarely absent). Nectariferous disc interstaminal and/or intrastaminal, or nectaries inserted on adaxial side of hypanthium, on ovary, on hypanthial base, as nectariferous hairs, or absent. Disc intrastaminal or absent.

Androecium Stamens usually five, 4+4 or 5+5 (sometimes three, 3+3, 6+6 to more than 100, rarely two), in one to five whorls. Filaments free from each other or more or less connate, free from tepals or adnate to petals (epipetalous). Anthers basifixed or dorsifixed, usually non-versatile, usually tetrasporangiate (sometimes disporangiate, thecae rarely up to more than 50), introrse, latrorse or extrorse, usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pore). Tapetum secretory. Staminodia absent or present (female flowers often with staminodia).

Pollen grains Microsporogenesis simultaneous. Pollen grains colpate, colporate or porate (sometimes pororate or stephanoporate) with three to 16 apertures, usually shed as monads (rarely tetrads), usually bicellular (sometimes tricellular) at dispersal. Exine tectate or semitectate, usually with columellate (sometimes granular or intermediate, acolumellate) infratectum, striate or reticulate, scabrate, echinate, spinulate or psilate.

Gynoecium Pistil composed of (one to) three or four (to six, rarely ten or twelve) usually more or less connate (rarely free) carpels. Ovary usually inferior (sometimes superior or semi-inferior), unilocular to quadrilocular (to sexalocular), sometimes with roof-like structure. Style single, simple, or stylodia (two or) three or four (to six), free or more or less connate; style often unifacial. Stigmas one or (two or) three (to five), commissural, often bilobate (in association with commissural lines), papillate or non-papillate, Dry or Wet type. Pistillodium absent.

Ovules Placentation axile, apical or (intrusively) parietal. Ovules one to more than 100 per carpel, anatropous, pendulous, horizontal or ascending, apotropous, usually bitegmic (rarely unitegmic), usually crassinucellar (sometimes tenuinucellar). Inner integument, when present, often delayed in development. Micropyle usually endostomal (sometimes bistomal, rarely absent). Nucellar cap or nucellar beak present. Hypostase present or absent. Megagametophyte usually monosporous, Polygonum type (rarely disporous, Allium type). Antipodal cells sometimes degenerating prior to fertilization, sometimes persistent, sometimes proliferating. Endosperm development ab initio nuclear. Endosperm haustorium chalazal or absent. Embryogenesis onagrad (sometimes caryophyllad?).

Fruit A loculicidal and/or septicidal capsule, a drupe, a berry, a schizocarp or a berry-like gourd (pepo, amphisarca?) with hard pericarp (rarely a samara or an assemblage of achenes).

Seeds Aril sometimes present. Operculum (formed by micropyle and hilum) often present. Seed coat testal, exotestal or mesotestal. Testa multiplicative, often vascularized, with lignified epidermis, unilayered or multilayered with sclerotic hypodermis and lignified inner layer. Exotesta usually thick. Endotesta usually membranous. Tegmen persistent, with tracheidal outer cells, or crushed or absent. Perisperm usually not developed (sometimes well developed, enclosing embryo). Endosperm rudimentary or absent. Embryo straight, usually well differentiated, without chlorophyll. Cotyledons two. Germination phanerocotylar or cryptocotylar.

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, bitter-tasting tetracyclic and pentacyclic triterpenes (cucurbitacins etc.), toxic sesquiterpenoid compounds, methylated and non-methylated ellagic acids, gallic acid, tannins, geraniins, karakin and other poisonous alkaloids, very poisonous bitter-tasting glycosides, triterpene saponins, cyclic polyvalent alcohols, punicic acid, eleostearic acid (isomere of punicic acid), special long-chain fatty acids (present in seed oils), myo-inisitol, and citrullin (α-amino-δ-ureidopentanoic acid, free amino acid) present. Cyanogenic compounds not found.

The crown clade of Cucurbitales, [[Coriariaceae+Corynocarpaceae]+[Cucurbitaceae+ [Tetramelaceae+[Datiscaceae+Begoniaceae]]]] has the potential synapomorphies (Stevens 2001 onwards): absence of uniseriate wood rays; filaments shorter than anthers in bud; anthers basifixed; absence of nectaries; and presence of free stylodia. Coriariaceae and Corynocarpaceae share the following synapomorphies: wood rays wide; sieve tube plastids without starch and protein inclusions; stomata paracytic; leaf margin entire; flowers small; hypogyny; sepals with imbricate quincuncial aestivation; petals thick, with broad base; carpels ascidiate; ovule one per carpel; vascular bundle extending into outer integument; cotyledons very large; and presence of ellagic acid.

The clade [Cucurbitaceae+[Tetramelaceae+[Datiscaceae+Begoniaceae]]] is characterized by the following features (Stevens 2001 onwards): flowers unisexual; perennial herbaceous growth; young stem with separate vascular bundles; absence of stipules; leaf margin serrate; medial vein ending in aggregation of translucent cells, lateral vein also entering tooth; carpels antesepalous or median carpel adaxial; roof-like structure present above ovary; stylodia marginal; stigmas large, elongated, bilobate; placentation parietal; ovules numerous per carpel; fruit many-seeded; presence of cucurbitacins; and absence of myricetin and ellagic acid.

The clade [Tetramelaceae+[Datiscaceae+Begoniaceae]] has the potential synapomorphies (Stevens 2001 onwards): pollen grains spheroidal; stigmas elongate; fruit an apically dehiscent septicidal capsule; testa with operculum formed by micropyle and hilum; exotestal cells honeycomb-shaped, inner walls strongly thickened and lignified; and cotyledons medium-sized. Datisca and Begoniaceae share the synapomorphies: herbaceous growth; and outer and inner integuments two cell layers thick. Tetramelaceae and Datiscaceae are both dioecious; have completely isomerous but not regularly pentamerous flowers (not in male plants of Datisca) with only small sepals and without petals (present in male plants of Octomeles).

Cucurbitaceae have the following features in common with [Tetramelaceae+[Datiscaceae+ Begoniaceae]] (Stevens 2001 onwards): herbaceous growth form; young stem with separate vascular bundles; leaf margin serrate, with cucurbitoid teeth (i.e. medial vein ending in pad of packed translucent cells, lateral veins also entering); absence of stipules; flowers unisexual; epigyny; carpels antesepalous or median carpel adaxial; extended roof-shaped structure on top of ovary (stylodia marginal), formed by ventral carpellary segments; stigmas large, elongated, bilobate; placentation parietal, often intrusive; ovules numerous per carpel; presence of cucurbitacins (oxidized triterpenes); and absence of myricetin and ellagic acid. The flowers are often androdioecious, the anthers basifixed and extrorse or latrorse, the free carpellary parts bifurcate, and the gynoecium trimerous.

Polygonanthaceae Croizat in Cact. Succ. J. (Los Angeles) 15: 64. Mai 1943; Anisophylleales (Benth. et Hook.f.) Takht. ex Reveal et Doweld in Novon 9: 550. 30 Dec 1999

Distribution Northern South America (Amazonas), tropical Africa, southern India, Sri Lanka, West Malesia.

Fossils Uncertain. Fossil pollen grains of Anisophylleaceae type are known from mid- and upper Miocene layers on Borneo. Fossil flowers of Platydiscus, a possible Anisophylleaceae (or Cunoniaceae?), have been found in upper Cretaceous strata in southern Sweden.

Habit Usually monoecious (sometimes polygamomonoecious or bisexual; in Combretocarpus dioecious), evergreen trees or shrubs. Most species are anisophyllous (Combretocarpus is isophyllous).

Vegetative anatomy Phellogen? Cambial stratification? Vessel elements with simple perforation plates; lateral pits alternate, simple or bordered pits. Imperforate tracheary xylem elements fibres with bordered pits, non-septate (also vasicentric tracheids). Wood rays multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal banded, aliform, lozenge-aliform or winged-aliform. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Parenchyma in Poga with lysigenous secretory cavities. Laticifers absent. Heartwood with resinous substances. Prismatic calciumoxalate crystals frequent.

Trichomes Hairs usually unicellular (sometimes multicellular, occasionally peltate), uniseriate, simple; glandular hairs usually absent (present near leaf axils in Anisophyllea disticha).

Leaves Alternate (spiral or distichous; in Anisophyllea tetrastichous), simple, entire, coriaceous, with ? ptyxis, usually dimorphic (in Combretocarpus monomorphic). Stipules usually absent (rarely two to four, very small, arising from petiole base, colleters?); leaf sheath absent. Petiole vascular bundle transection simple. Leaf base in Anisophyllea asymmetrical. Venation usually palmate (in Combretocarpus pinnate). Stomata usually paracytic. Cuticular wax crystalloids as platelets (in Polygonanthus cupular). Leaf margin entire.

Inflorescence Axillary, usually panicle, raceme or spike (flowers rarely solitary).

Flowers Actinomorphic, usually small. Epigyny. Sepals (three or) four (to 16), in one whorl, with valvate aestivation, persistent, with mucilaginous inner epidermal walls, postgenitally coherent. Petals (three or) four (to 16; in Combretocarpus often absent), with valvate or open aestivation, clawed, usually trilobate, quinquelobate or septalobate (in Poga entire), more or less enclosing groups of free stamens. Nectariferous disc crenate, usually both interstaminal and intrastaminal (nectariferous disc in Combretocarpus only intrastaminal; in Anisophyllea formed by protrusions between and behind each stamen).

Androecium Stamens 3+3, 4+4 or 5+5, obdiplostemonous. Filaments narrow, free from each other and from tepals, inflexed in bud. Anthers dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia present or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate to tricolporoidate (in Anisophyllea usually dicolpate), shed as monads or tetrads (or polyads?), bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, usually reticulate or punctate (sometimes striate or smooth).

Gynoecium Pistil composed of (three or) four (or five) carpels connate in lower part. Ovary inferior, (trilocular or) quadrilocular (or quinquelocular); sometimes with roof-like structure above ovary. Stylodia (three or) four (or five), free, sometimes hollow; compitum absent. Stigmas widened or punctate, type? Pistillodium often present in male flowers.

Ovules Placentation apical-axile or parietal. Ovule usually one per carpel, anatropous, pendulous, epitropous, bitegmic or unitegmic (Anisophyllea, Combretocarpus), crassinucellar. Micropyle bistomal (Poga, Polygonanthum; in Anisophyllea slit-shaped). Outer integument four or five cell layers thick. Inner integument approx. two cell layers thick. Integument in Anisophyllea seven to nine cell layers thick (unitegmic ovules). Parietal tissue approx. one cell layer thick. Nucellar cap present. Megagametophyte usually monosporous, Polygonum type (in Combretocarpus disporous, 8-nucleate, Allium type). Antipodal cells degenerating prior to fertilization (mature megagametophyte quinquecellular). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit Usually a drupe (rarely a capsule; in Combretocarpus a samara), sometimes with accrescent sepals.

Seeds Aril present or absent. Seed coat testal. Testa usually multiplicative, sometimes vascularized, approx. ten to c. 30 cell layers thick, thin and one-layered (Combretocarpus), or thick and multilayered (Anisophyllea, Poga). Outer exotestal epidermis cells cuboid, with thick, lignified walls. Mesotestal cell walls sometimes lignified. Tegmen crushed or absent. Exotegmen not sclerified. Perisperm not developed. Endosperm absent. Embryo fusiform, with long hypocotyl, chlorophyll? Cotyledons two, usually indistinct or absent (in Combretocarpus small). Germination cryptocotylar.

Phytochemistry Very insufficiently known. Tannins present. Alkaloids and saponins not found. Aluminium accumulated.

Use Timber (Combretocarpus rotundatus, Poga oleosa), seed oils (Poga).

Distribution Tropical East Africa, southeastern Turkey to northern Iran, southwestern Australia, southern California, Mexico, Central America, the West Indies, South America south to central Argentina.

Habit Usually monoecious or dioecious (rarely bisexual), achlorophyllous herbaceous endophytes without rhizome or normal roots. Root or stem holoendoparasites (Apodanthes on Salicaceae [Casearia, Xylosma]; Pilostyles on Papilionoideae in Fabaceae).

Vegetative anatomy Mycorrhiza absent. Hypha-like cell-threads invading host plants, forming endophytic system inside host roots. Phellogen absent. Secondary lateral growth absent. Vessels absent. Imperforate tracheary elements? Wood rays absent. Axial parenchyma? Stomata anomocytic. Sieve tube plastids S₀ type, without starch or protein inclusions. Cells with calciumoxalate crystals. Oils or mucilage absent.

Trichomes Hairs (in flowers) unicellular or multicellular, uniseriate; vesicular hairs present.

Flowers Actinomorphic, small, usually with evil smell. Epigyny or half epigyny. Floral (tepaloid) scales with entire margin probably in three di-, tri-, tetra- or hexamerous whorls of eight to 15 scales in total (2+4+4 or 3+6+6, etc.), at least inner scales probably tepals and outer scales bracts, usually with open or imbricate (sometimes contorted) aestivation, free or connate at base, usually persistent (outer scales in Apodanthes caducous); inner whorl of scales with adaxial hair tufts. Thin tissue, diaphragma, and special outgrowth, ramenta, inserted on top of perigonal tube. Gynostemium present. Nectariferous disc somewhat quadrangular or sexangular, extrastaminal (formed by base of innermost scales), in Apodanthes and Pilostyles with stomata. Ring of vesicular hairs present in male flowers on upper part of pistillodium, immediately above synandrium.

Androecium Stamens congenitally connate into synandrium, not differentiated into filaments and anthers. Thecae five to more than 50, in (one or) two or three (or four) whorls, extrorse, longicidal (in Pilostyles dehiscing by longitudinal slits, sometimes between thecal whorls, and in Apodanthes by longitudinal slits) or poricidal (dehiscing by apical pore). Endothecium absent. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains monoporate, monocolpate, triporate, tricolpate (Apodanthes, Pilostyles), or inaperturate (Pilostyles), shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, psilate.

Gynoecium Pistil usually composed of four (or five) distally connate antepetalous carpels. Ovary inferior or semi-inferior, unilocular. Style single, simple, very short, hollow, with H-shaped (in transverse view) stylar canal, compitum; nectary present on stylar base. Stigma single, simple (Apodanthes, Pilostyles) or somewhat quadrilobate (Pilostyles), annular, hemispherical, relatively large, inserted below apex of central column, papillate, Wet type. Male flowers with pistillodium with basal nectary and vesicular hairs at least on margin.

Ovules Placentation parietal. Ovules c. 15 to more than 90 per carpel, anatropous, bitegmic, tenuinucellar. Micropyle bistomal or endostomal (Pilostyles), or absent (Apodanthes, Pilostyles). Outer integument two cell layers thick. Inner integument two cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Seeds Aril absent. Elaiosome present, formed from exotesta. Testa thin-walled, mucilaginous, exotestal. Exotestal cells? Endotesta? Exo(?)tegmic cell walls strongly lignified. Endotegmen? Perisperm not developed. Endosperm copious. Embryo rudimentary, undifferentiated, without chlorophyll? Cotyledons two? Germination?

Begoniales Bercht. et J. Presl, Přir. Rostlin: 270. Jan-Apr 1820 [‘Begoniae’]; Begoniineae Engl., Syllabus, ed. 2: 156. Mai 1898; Begonianae Doweld, Tent. Syst. Plant. Vasc.: xxxix. 23 Dec 2001

Distribution Tropical and subtropical regions on both hemispheres, with their highest diversity in tropical Asia and northern South America.

Habit Usually monoecious (male flowers are produced ab initio, and subsequently female flowers; rarely dioecious), usually more or less succulent perennial herbs with tuber (in Hillebrandia round) or rhizome (in Begonia rhizome, rarely tuber), sometimes climbing, sometimes somewhat woody below and frutescent. Nodes swollen.

Vegetative anatomy Phellogen ab initio subepidermal. Medulla sometimes storied by diaphragms. Cortical and medullary vascular bundles usually present. Primary vascular tissue cylinder of vascular bundles. Pericyclic envelope absent. Vessel elements with simple or scalariform perforation plates; lateral pits scalariform, simple pits. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, non-septate. Wood rays multiseriate, heterocellular, or absent. Axial parenchyma paratracheal scanty. Fibres sometimes storied. Tyloses often abundant. Sieve tube plastids S type. Nodes 3:3, trilacunar with three leaf traces, or 5:5, pentalacunar with five traces. Brachysclereids and non-calcified cystoliths present. Calciumoxalate as prismatic crystals, druses and raphides.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple or branched, sometimes bristle-like (setular), candelabra-shaped or stellate, often large and flat; peltate-lepidote glands and sessile spherical pearl glands present.

Leaves Usually alternate (usually distichous, sometimes spiral, rarely opposite), usually simple (rarely palmately compound), entire or palmately lobed, with usually conduplicate (vertically or laterally; sometimes supervolute-curved) ptyxis. Stipules usually large, cauline-extrapetiolar or intrapetiolar, caducous or persistent; leaf sheath absent. Petiole vascular bundle transection annular; petiole sometimes with central vascular bundles. Leaf base usually strongly asymmetrical. Venation palmate. Stomata helicocytic with three to six subsidiary cells (often in two whorls) or anisocytic (rarely paracytic or diacytic?), often only on adaxial side of lamina. Cuticular wax crystalloids? Mesophyll with or without sclerenchymatous idioblasts. Cystoliths (non-calcified) usually present. Hydathodes usually absent. Leaf margin usually serrate or crenate (rarely entire); leaf teeth begonioid, supplied by several veins; medial vein ending in aggregation of translucent cells, lateral vein dominant, also entering tooth. Caducous pearl glands present.

Inflorescence Usually axillary (in Begonia rarely epiphyllous), dichasial or bostrycoid. Female flowers in Begonia usually developed in cymose inflorescence after male flower anthesis (inflorescence in Symbegonia subclade racemose and female flowers produced prior to male flowers).

Flowers More or less zygomorphic or bisymmetrical. Usually epigyny (in Hillebrandia half epigyny). Sepals usually two, with valvate or imbricate aestivation, petaloid (rarely six to eight; in Hillebrandia five, larger to much larger than petals), usually free (sometimes connate into tube). Petals usually two, with valvate aestivation, or absent (in Hillebrandia five, smaller to much smaller than sepals and possibly staminodia; in female flowers of Begonia two to nine; tepals in male flowers of Begonia usually two or four, sometimes three or five to eight), usually free (rarely connate into campanulate corolla). Nectary usually absent (nectaries rarely present at stylodium bases). Disc absent.

Androecium Stamens three to c. 50 to more than 100, in two to five whorls, centrifugally developing (in Hillebrandia with branched vascular bundles). Filaments free or more or less connate, free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, extrorse or latrorse, usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores), yellow; connective often exlpanded. Tapetum secretory. Female flowers often with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolp(or)ate, shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, often striate.

Gynoecium Pistil composed of (two or) three (or six; in Hillebrandia five; in Begonia sometimes one) carpels connate in lower part. Ovary usually inferior (in Hillebrandia semi-inferior), unilocular (in Hillebrandia and some species of Begonia) or secondarily usually trilocular (rarely bilocular or sexalocular), usually three-winged; carpels strongly developed ventrally forming roof-like structure above locule, this forming perianth tube base; calyx tube with widely separated stylodia inserted on roof edge. Stylodia (two or) three (or six), usually free (sometimes connate at base; in Hillebrandia peripheral; in Begonia central), short, usually deeply bifid. Nectaries inserted at stylodium bases in some ornithophilous species. Stigmas often twisted (yellow, resembling anthers), papillate, Dry type. Pistillodium absent.

Ovules Placentation usually axile (in Hillebrandia axile at base and parietal at apex; in Begonia axile to parietal), with large usually bilamellate placentae. Ovules numerous per carpel, anatropous, bitegmic, crassinucellar. Micropyle bistomal, Z-shaped (zig-zag). Outer integument ? cell layers thick. Inner integument approx. two cell layers thick. Endothelium present. Parietal tissue approx. two cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad.

Fruit Usually a loculicidal capsule (in Hillebrandia septicidal, dehiscing at apex between stylodia; in Begonia rarely also septicidal), usually winged (often asymmetrically winged; rarely a berry).

Seeds Aril absent. Funicle surrounded by collar formed as extension of testa. Operculum formed by micropyle and hilum, and surrounding annulus of collar cells. Seed coat exotestal. Exotestal cells with honeycomb-like arrangement, with heavily thickened and lignified inner walls? Endotesta absent or almost absent Tegmen absent or almost absent. Perisperm not developed. Endosperm sparse or absent. Embryo small, straight, little to well differentiated, chlorophyll? Cotyledons two. Germination phanerocotylar.

Phytochemistry Flavonols (quercetin), cyanidin, cucurbitacins, tannins, calcium oxalate, oxalic acid, proanthocyanidin, saponins and cyanidin glycosides present. Ellagic acid, myricetin, alkaloids, and cyanogenic compounds not found.

Coriariales Lindl., Nix. Plant.: 11. 17 Sep 1833 [‘Coriales’]; Coriariopsida Parl., Fl. Ital. 5: 486. 1872 [’Coriarineae’]; Coriariineae Engl., Syllabus, ed. 2: 142. Mai 1898

Distribution Western Mediterranean, the Himalayas to Japan, Taiwan, the Philippines, New Guinea, New Zealand, islands in southwestern Pacific, tropical America.

Fossils Macrofossils assigned to Coriaria longaeva are known from Oligocene layers in France, and seeds and pollen grains of Coriariaceae have been described from the Miocene of several European sites.

Habit Usually dioecious (sometimes monoecious, andromonoecious, gynomonoecious or polygamomonoecious, rarely bisexual), evergreen or deciduous shrubs (sometimes suffrutices or small trees). Some branches with limitied growth similar to pinnate leaves. Buds usually perulate.

Vegetative anatomy Root nodules containing nitrogen-fixing endosymbiotic actinobacteria (Frankia) present in most species. Phellogen ab initio superficial. Primary medullary strands wide. Vessel elements present in multiples. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements tracheids or libriform fibres with simple or bordered pits, non-septate (also vasicentric tracheids). Wood rays multiseriate, heterocellular. Axial parenchyma paratracheal scanty, confluent or vasicentric. Wood elements partially storied. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Prismatic calciumoxalate crystals abundant.

Leaves Usually opposite (sometimes verticillate, rarely alternate spiral), simple, entire, coriaceous, with flat ptyxis. Stipules minute, caducous; leaf sheath absent. Petiole vascular bundle transection arcuate. Venation palmate. Stomata paracytic. Cuticular waxes absent. Epidermis with secretory cells. Calciumoxalate crystals abundant. Tannins very abundant. Leaf margin entire or weakly dentate.

Flowers Actinomorphic, small. Hypogyny. Sepals five (or six), with imbricate quincuncial aestivation, persistent, free. Petals five (or six), with valvate or open aestivation, persistent, free, usually adaxially keeled. Nectary absent. Disc absent.

Androecium Stamens usually 5+5 (rarely 6+6), diplostemonous. Filaments thin, free from each other, free from tepals or antepetalous filaments adnate to petal keel. Anthers basifixed or slightly dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective narrow. Tapetum secretory, with binucleate to quadrinucleate cells. Female flowers often with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–4)-colporate or 3(–4)-pororate, starchy, shed as monads, usually tricellular (sometimes bicellular) at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Carpels usually five, antesepalous (rarely ten or twelve), free from each other or connate at base (although bulging and seemingly free); carpel ascidiate? Ovary superior, unilocular (apocarpy). Stylodia usually five, filiform, subbasal; compitum present. Stigmatic areas surrounding style, papillate, Dry type. Male flowers often with pistillodium.

Ovules Placentation apical-axile. Ovule one per carpel, anatropous, pendulous, apotropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument three or four cell layers thick. Inner integument two or three cell layers thick. Parietal tissue approx. eight cell layers thick. Nucellar cap approx. four cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit: A schizocarp with nutlike mericarps or an assemblage of achenes with persistent subbasal stylodia and enclosed by carnose accrescent petals, resulting in baccate or drupaceous appearance.

Seeds Aril absent. Seed coat exotestal. Exotesta consisting of cuboid, tanniniferous cells with thick lignified? walls. Endotesta and tegmen indistinct. Perisperm not developed. Endosperm sparse, oily, or absent. Embryo straight, oily, chlorophyll? Cotyledons two, large. Germination phanerocotylar.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), toxic sesquiterpenoid compounds, methylated and non-methylated ellagic acids, gallic acid, tannins, alkaloids, geraniins, cyclic polyvalent alcohols, coriolic fatty acid (in seeds), and myo-inisitol present. Triterpene saponins? Proanthocyanidins and cyanogenic compounds not found.

Engler in Engler et Prantl, Nat. Pflanzenfam., Nachtr. 1: 215. Oct 1897, nom. cons.

Corynocarpales Takht., Divers. Classif. Fl. Pl.: 340. 24 Apr 1997; Corynocarpanae Takht., Divers. Classif. Fl. Pl.: 340. 24 Apr 1997

Distribution The Aru Islands, New Guinea, New Britain, New Ireland, northeastern and eastern Australia, Melanesia, New Zealand.

Vegetative anatomy Phellogen ab initio subepidermal. Young stem with separate vascular bundles. Cambium storied. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, non-septate. Wood rays multiseriate, homocellular or heterocellular, very wide. Wood elements partially storied. Axial parenchyma paratracheal scanty, confluent, vasicentric or wide-banded. Sieve tube plastids S type. Nodes 3:3?, trilacunar with three? leaf traces. Prismatic calciumoxalate crystals abundant.

Leaves Leaves alternate (spiral), simple, entire, coriaceous, with conduplicate ptyxis. Stipule single, intrapetiolar, caducous, or absent; leaf sheath absent. Petiole vascular bundle transection forming line. Venation pinnate, brochidodromous. Stomata paracytic. Cuticular wax crystalloids? Leaf margin entire.

Flowers Actinomorphic, small. Hypanthium short. Hypogyny. Sepals five, with imbricate aestivation, free. Petals five, with imbricate aestivation, free. Nectaries five, intrastaminal, staminodial. Disc consisting of five separate parts.

Androecium Fertile stamens five, obhaplostemonous, alternisepalous, antepetalous. Filaments free, adnate at base to petals, inflexed in bud. Anthers dorsifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia five, petaloid, extrastaminal or interstaminal, alternating with stamens, fimbriate, each with basal adaxial antesepalous nectary.

Pollen grains Microsporogenesis simultaneous? Pollen grains dicolp(or)ate, heteropolar (one short colpus at each pole), shed as monads, ?-cellular at dispersal. Exine tectate, with granular to intermediate infratectum, scabrate to psilate.

Gynoecium Pistil composed of two connate carpels, one of which more or less reduced and sterile leading to pseudomonomery. Ovary superior, unilocular or bilocular. Stylodium usually single, simple, short, conduplicate (stylodia rarely two). Stigma capitate, Dry type. Pistillodium absent.

Ovules Placentation apical. Ovule one per fertile carpel, anatropous, pendulous, apotropous or epitropous?, with erect micropyle, bitegmic, crassinucellar. Micropyle endostomal. Outer integument c. 11 (to c. 30?) cell layers thick. Inner integument two or three cell layers thick. Megagametophyte monosporous, Polygonum type. Antipodal cells proliferating (with up to eight cells). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis caryophyllad?

Seeds Aril absent. Testa pachychalazal?, ab initio thick, vascularized, finally crushed. Tegmen? Perisperm not developed. Endosperm sparse, starchy, or absent. Embryo straight, well differentiated, oily and starchy, chlorophyll? Cotyledons two, large, thick. Germination phanerocotylar.

Phytochemistry Flavonols (kaempferol), ellagic acid, very poisonous bitter glucosides (at least in bark and seeds), and karakin (in fruits) present. Proanthocyanidins, saponin, and cyanogenic compounds not found.

Nhandirobaceae T. Lestib., Botanogr. Elém.: 515. Jun 1826 [’Nandhirobées’]; Zanoniaceae Dumort., Anal. Fam. Plant.: 28, 29. 1829; Bryoniaceae G. Mey., Chloris Han.: 6, 112. Jul-Aug 1836; Cyclantheraceae Lilja, Skånes Fl., ed. 2: 716, 980. Apr-Dec 1870 [’Cyclanthereae’]; Cucurbitineae Engl., Syllabus, ed. 2: 190. Mai 1898

Distribution Tropical and subtropical regions, with their largest diversity in South American rainforests and arid regions in Africa, few species in Australasia and in temperate regions.

Fossils Seeds have been described from the Paleocene and the Eocene of England. Fossil hexacolpate or stephanocolpate pollen grains, Hexacolpites echinatus, have been described from the Oligocene of Cameroon.

Habit Monoecious, andromonoecious, gynomonoecious, polygamomonoecious, dioecious, androdioecious, and gynodioecious (in Actinostemma and Schizopepon sometimes bisexual), usually perennial herbs, mostly climbing or winding (rarely lianas, shrubs or tree, secondarily woody, or annual herbs; Dendrosicyos extremely pachycaul and secondarily arborescent with soft juicy stem). Many species are xerophytes. In leaf axils usually lateral and often branched tendrils formed by modified lateral shoots with lower part corresponding to stem branch and climbing upper part corresponding to strongly modified leaves (tendrils sometimes modified into spines [i.a. in Acanthosicyos, Momordica] or absent). Tendrils simple or branched; branched tendrils with or without sensitive coiling base (zanonioid tendrils), sometimes with terminal adhesive pads. Tendrils often part of axillary complex together with axillary buds and inflorescences. Many species provided with epigeal tuberous pachypodium (root stock) formed by swollen hypocotyl (cauduciform succulents).

Vegetative anatomy Root phellogen superficial. Stem phellogen ab initio superficially to deeply seated. Primary vascular tissue with separate bundles or consisting of cylinder of bundles. Young stem usually with bicollateral vascular bundles, often arranged in two cylinders. Successive cambia present in at least Bryonia and Ecballium. Cortical vascular bundles usually present. Cortex with sclerenchymatous envelope. Pericyclic sheath absent. Secondary lateral growth normal or anomalous (via concentric cambia or from simple cambial cylinder). Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, septate (also vasicentric tracheids). Wood rays multiseriate, heterocellular. Axial parenchyma apotracheal or paratracheal aliform or banded. Wood elements often storied. Tyloses often frequent. Extrafascicular phloem usually present in cortex outside sclerenchymatous envelope (in species with bicollateral vascular bundles). Sieve tube plastids S type. Nodes 3:3, trilacunar with three leaf traces. Branched idioblasts with bitter substances. Cystoliths usually present. Raphids present in at least Cucurbita pepo and Ecballium; calcium oxalate crystals and cystoliths, calcium carbonate bodies of varying shape (in some genera), crystalline silica grains (in at least Cucurbita).

Trichomes Hairs unicellular, simple, verrucate or often as prickles; glandular hairs often frequent; cell walls often calcified; calcified cystoliths and similar structures often present at hair bases or in adjacent cells.

Leaves Alternate (spiral), palmately compound or simple and usually palmately lobed (sometimes entire), with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate or as ring of arcuate bundles, unequal in size (larger bundles bicollateral); leaf supplied by one vascular strand from outer ring of bundles and by branches from two additional strands of outer bundle ring. Pericyclic envelope usually absent. Extrafloral nectaries frequent on abaxial surface of lamina and/or sometimes on petiole, node or leaf margin. Venation usually palmate. Stomata usually anomocytic. Cuticular wax crystalloids usually absent, sometimes as rosettes of platelets (Fabales type), or as irregular platelets or terete rodlets, chemically dominated by triterpenoids, etc. Hydathodes often present. Leaf margin usually serrate (rarely entire); leaf teeth often cucurbitoid.

Inflorescence Axillary, cymose, umbel, raceme, fascicle, pseudopanicle, or flowers solitary. Female flowers often solitary in leaf axils on main branch, male inflorescence arising from this first node. Floral prophylls (bracteoles) often absent. Floral bracts sometimes with extrafloral nectaries.

Flowers Usually actinomorphic (sometimes with zygomorphic androecium), often large. Hypanthium, formed by lower parts of perianth and stamens, usually present. Usually epigyny (rarely half epigyny). Sepals (three to) five (to seven), with imbricate or open aestivation, usually more or less connate (rarely absent). Petals (three to) five (to seven), with valvate, induplicate-valvate or imbricate quincuncial aestivation, in lower parts usually connate into campanulate perigonal tube. Nectary usually disc-shaped (sometimes covered by tissue lobe). Male flowers usually with nectaries on adaxial side of hypanthium. Female flowers with nectaries on ovary, on hypanthial base, or as nectariferous glandular hairs. Oil-secreting hairs sometimes present. Disc intrastaminal or absent.

Androecium Stamens usually five (sometimes three; in, e.g., Gurania, Helmontia and Psiguria two), haplostemonous, antesepalous, alternipetalous. Filaments usually entirely or almost entirely connate in fascicles 2+2+1 or all five stamens into central columella (sometimes free or five free stamens or three free filaments due to fusion of two adjacent stamens, although still five anthers due to incomplete fusion of thecae; in Fevillea five free stamens with bilocular anthers), often adnate to petals, usually inserted into hypanthium. Anthers dorsifixed, non-versatile, straight or arcuate to triplicate or (often strongly) sigmoid, often connate into tube (in Thladiantha connate 4+1), when three anthers then usually two tetrasporangiate and one disporangiate (monothecal), when five anthers then all disporangiate (monothecal), extrorse, longicidal (dehiscing by longitudinal slits); connective sometimes prolonged at apex. Tapetum secretory. Female flowers with rudimentary staminodia (male flowers in Gerrardanthus with one out of five stamens modified into staminodium).

Pollen grains Microsporogenesis simultaneous. Pollen grains colpate, colporate or porate (sometimes stephanoporate) with three to 16 apertures, starchy, sometimes operculate, sometimes very large (40–70 µm or more), usually shed as monads (in Gurania and Psiguria tetrads), bicellular at dispersal. Exine tectate to semitectate, with columellate or acolumellate infratectum, perforate or reticulate, echinate, spinulate, striate or microstriate. Pollen grain germinating with several pollen tubes (polysiphony).

Gynoecium Pistil composed of usually (two or) three (to five) connate carpels (in Sicyoeae often one carpel); median carpel sometimes abaxial. Ovary usually inferior (rarely semi-inferior), unilocular, although usually almost filled up by enlarged placentae, sometimes fused in centre and seemingly multilocular; carpels strongly developed ventrally forming roof-like structure above locule, this forming perianth tube base; calyx tube with widely separated stylodia inserted on roof edge. Stylodia (two or) three (to five), entirely (sometimes only below) connate (style simple) or more or less separate. Stigmas one or (two or) three (to five), commissural, often bilobate (in connection with commissural lines), papillate or non-papillate, Dry or Wet type. Pistillodium absent (small processes alternating with stamens perhaps representing reduced carpels).

Ovules Placentation intrusively parietal (sometimes seemingly axile). Ovules usually numerous (sometimes few; in Sicyoeae one) per carpel, anatropous, pendulous or horizontal to ascending, bitegmic, crassinucellar. Micropyle usually endostomal (rarely bistomal). Outer integument four to eight cell layers thick, vascularized. Inner integument two or three (to at least six) cell layers thick. Parietal tissue five to eleven cell layers thick. Nucellar cap and nucellar beak present. Hypostase present or absent. Megagametophyte usually monosporous, Polygonum type (rarely disporous, 8-nucleate, Allium type), sometimes extremely long (cf Santalales). Antipodal cells usually degenerating. Endosperm development ab initio nuclear. Chalazal end of megagametophyte usually forming tubular endosperm haustorium. Embryogenesis onagrad.

Fruit Usually a single- to many-seeded berry or berry-like gourd (pepo, amphisarca?) with hard pericarp (rarely a dry och fleshy capsule, a samara or a fleshy explosion fruit; fruit in Actinostemma and Bolbostemma operculate).

Seeds Aril absent; arilloid (derived from nearby carpellary tissue) often present. Seeds often flattened, sometimes winged. Operculum? Seed coat mesotestal. Testa multiplicative, vascularized, often complex, with lignified epidermis, unilayered or multilayered sclerotic hypodermis (absent in Coniandreae) and finally lignified inner layer, with single cell layer usually rich in asterosclereids (sometimes osteosclereids), sclereid layers usually strongly separated from other cells; testa often with aerenchyma inside epidermis, thick-walled and lignified, but little differentiated from hypodermal layer; inner sclerenchymatous layers often with anticlinal cell divisions, brachysclereid. Exotesta usually thick, often palisade or cuboid (in some species with multilayered epidermis). Endotesta usually membranous. Tegmen persistent, with tracheidal outer cells. Perisperm usually not developed (sometimes well developed, enclosing embryo). Endosperm rudimentary or absent. Embryo straight, well differentiated (plumule often with distinct leaves), without chlorophyll; often with crystalloid inclusions in protein bodies. Cotyledons two, large, flat. Germination phanerocotylar or cryptocotylar. Seedlings often with cortical outgrowth on lower side of axis in transition zone between root and stem.

Cytology n = 7–24 (9–14, 16, 20, 22, 33, 44, 66, 88); x = 12 – Fixed polyploidy (n = 20) occurring in Cucurbiteae.

DNA Plastid gene infA lost/defunct (Luffa). Inversion of 35–40 bp present in plastid trnL/trnF-intergenic spacer (absent in other Cucurbitales; absent in Neoalsomitra and Cucurbita digitata). Cucurbitaceae usually with labile genome organization: two (or three) cases (out of extremely few known among angiosperms) of independent transfer of gene rbcL from plastid to mitochondrial genome (Cucumis sativus, Cucurbita maxima and Cucurbita pepo). Mitochondrial coxI intron present in at least Citrullus, Cucumis and Melothria. Cucumis with largest known mitochondrial genome among angiosperms (1.500 kb in C. sativus, 2.400 kb in C. melo).

Phytochemistry Flavonols (kaempferol, quercetin), extremely bitter-tasting tetra- and pentacyclic triterpenoids (cucurbitacins etc.), alkaloids, triterpene saponins, punicic acid (C₁₈H₃₀O₂), eleostearic acid (isomere of punicic acid, in Joliffieae), long-chain fatty acids (in seed oils) and citrullin (free non-protein amino acid α-amino-δ-ureidopentanoic acid) present. Ellagic acid, tannins, and proanthocyanidins not found.

Use Ornamental plants, fruits, vegetables, water sources, medicinal plants, containers and musical instruments (Lagenaria siceraria).

Systematics A probable topology of Cucurbitaceae is (Schaefer & Renner 2011): [Gomphogyneae+[Alsomitra macrocarpa+[[Triceratieae+Zanonieae]+[Actinostemmateae+[Indofevilleeae+[Thladiantheae+[Siraitieae+[Momordiceae+[Joliffieae+[Bryonieae+[Sicyoeae+[Schizopeponeae+[Coniandreae+[Benincaseae+Cucurbiteae]]]]]]]]]]]]]].

Gomphogyneae Benth. et Hook. f., Gen. Plant. 1: 820. Sep 1867

5/56. Bayabusua (1; B. clarkei; the Malay Peninsula), Gomphogyne (6; G. bonii, G. cirromitrata, G. cissiformis, G. heterosperma, G. nepalensis, G. peekelii; eastern Himalayas to central China and Southeast Asia), Gynostemma (c 20; East and tropical Asia to New Guinea), Hemsleya (27; the Himalayas, East Asia, Indochina, East Malesia), Neoalsomitra (2; N. clavigera, N. sarcophylla; tropical Asia to eastern Queensland and Fiji). – East and tropical Asia, eastern Queensland, Fiji, with their largest diversity in Southeast Asia. Lianas. Tendrils usually apically bifid, often with adhesive pads. Petals with multicellular nectariferous hairs. Stamens three or five, largely connate. Filaments adnate at base to petals. Anthers dithecal and/or monothecal. Fruit a capsule or berry. n = 11 (Gynostemma), 14 (Hemsleya), 16 (Gomphogyne).

[Alsomitra macrocarpa+[[Triceratieae+Zanonieae]+[Actinostemmateae+[Indofevilleeae+[Thladiantheae+[Siraitieae+[Momordiceae+[Joliffieae+[Bryonieae+[Sicyoeae+[Schizopeponeae+[Coniandreae+[Benincaseae+Cucurbiteae]]]]]]]]]]]]]

Alsomitra?

1/1. (1; A. macrocarpa; Southeast Asia, Malesia to New Guinea). – Liana. n = ? – Alsomitra has sometimes been identified as sister to the remaining Cucurbitaceae, yet this may be the result of long-branch attraction.

[[Triceratieae+Zanonieae]+[Actinostemmateae+[Indofevilleeae+[Thladiantheae+[Siraitieae+[Momordiceae+[Joliffieae+[Bryonieae+[Sicyoeae+[Schizopeponeae+[Coniandreae+[Benincaseae+Cucurbiteae]]]]]]]]]]]]

[Triceratieae+Zanonieae]

Triceratieae A. Rich. in R. de la Sagra, Hist. Fis. Cuba, Bot. 10: 298. 1845 [’Triceratiae’]

5/23. Fevillea (7; F. anomalosperma, F. bahiensis, F. cordifolia, F. narae, F. pedatifolia, F. pergamentacea, F. trilobata; southern Mexico, Central America, tropical South America), Anisosperma (1; A. passiflora; Brazil), Pteropepon (4; P. argentinense, P. deltoideus, P. oleiferum, P. parodii; Brazil, Peru, northern Argentina), Cyclantheropsis (3; C. madagascariensis, C. occidentalis, C. parviflora; tropical Africa, Madagascar), Sicydium (8; tropical America). – Tropical Africa, Madagascar, tropical America. Lianas. Tendrils simple or apically bifid. Petals with multicellular nectariferous hairs? Stamens one to five. Filaments adnate at base to petals. Anthers dithecal or monothecal. Female flowers with five staminodia. Fruit a pepo, samara or achene. n = ?

Zanonieae Benth. et Hook. f., Gen. Plant. 1: 820. Sep 1867

4/11. Gerrardanthus (5; G. grandiflorus, G. lobatus, G. macrorhizus, G. paniculatus, G. tomentosus; tropical and southern Africa), Siolmatra (2; S. brasiliensis, S. pentaphylla; Amazonas), Zanonia (1; Z. indica; tropical Asia), Xerosicyos (3; X. danguyi, X. perrieri, X. pubescens; Madagascar). – Tropical and southern Africa, Madagascar, tropical Asia to New Guinea, the Amazon Basin. Lianas or twining herbs. Xerosicyos comprises twining woody leaf succulents. Tendrils usually bifid. Inflorescence with internode below leaf under first flower. Petals in Xerosicyos free. Petals with multicellular nectariferous hairs. Stamens (four or) five. Filaments adnate in lower parts to petals. Anthers monothecal. Fruit a capsule. n = ?

[Actinostemmateae+[Indofevilleeae+[Thladiantheae+[Siraitieae+[Momordiceae+[Joliffieae+[Bryonieae+[Sicyoeae+[Schizopeponeae+[Coniandreae+[Benincaseae+Cucurbiteae]]]]]]]]]]]

Actinostemmateae H. Schaef. et S. S. Renner in Taxon 60: 130. 1 Feb 2011

1/4. Actinostemma (4; A. lobatum, A. paniculatum, A. parvifolium; two species in China; A. tenerum; India, China and eastern Siberia to the Korean Peninsula and Japan, Taiwan, Laos, Vietnam). – Twining herbs. Tendrils usually bifid. Petals with multicellular nectariferous hairs. Stamens five or 2+2+1. Filaments adnate at base to petals. Anthers monothecal. Style single. Ovule pendulous. Fruit a pyxidium. Testa often winged. n = 8.

[Indofevilleeae+[Thladiantheae+[Siraitieae+[Momordiceae+[Joliffieae+[Bryonieae+[Sicyoeae+[Schizopeponeae+[Coniandreae+[Benincaseae+Cucurbiteae]]]]]]]]]]

Usually only branches of tendrils coiled. Young stem with bicollateral vascular bundles. Extrafascicular phloem associated with adaxial phloem of vascular bundles. Axillary complex consisting of tendril, bud, male inflorescence, and female flower (collateral). Nectary usually disc-shaped, parenchymatous, with stomata. Male flowers with well developed hypanthium. Style single. Secretion-filled stylar canal present. Ovules horizontal to erect. Outer integument six to ten cell layers thick. Inner integument (one or) two to five (or six) cell layers thick. Parietal tissue five to eleven cell layers thick. Antipodal cells degenerating. Fruit usually baccate. Testa not winged. Exotesta enlarged, palisade or cuboid, mucilaginous, with distinctive sclereid layer and often strongly thickened cell walls. Innermost layer chlorenchymatous. Non-protein amino acids present.

Indofevilleeae H. Schaef. et S. S. Renner in Taxon 60: 130. 1 Feb 2011

1/1. Indofevillea (1; I. khasiana; Assam, Bhutan, Tibet). – Liana. Tendrils zanonioid, apically bifid. Stamens 2+2+1, adnate at base to corolla tube. Anthers monothecal. Fruits dry, indehiscent, with thick lignified pericarp. n = ?

[Thladiantheae+[Siraitieae+[Momordiceae+[Joliffieae+[Bryonieae+[Sicyoeae+[Schizopeponeae+[Coniandreae+[Benincaseae+Cucurbiteae]]]]]]]]]

Thladiantheae H. Schaef. et S. S. Renner in Taxon 60: 130. 1 Feb 2011

2/30–35. Baijiania (5; B. borneensis, B. decipiens, B. smitinandii, B. taiwaniana, B. yunnanensis; southern China, Thailand, Laos, Borneo, Taiwan), Thladiantha (25–30; southern Russia and western Asia, India, the Himalayas, Tibet, China, the Korean Peninsula, Japan, Taiwan, Southeast Asia, Malesia to New Guinea). – Temperate Asia, East and tropical Asia to Taiwan and Malesia. Lianas or twining herbs. Tendrils simple or bifid. Stamens 2+2+1, adnate to corolla tube (epipetalous). Anthers monothecal. Fruits baccate. n = 9 (Thladiantha), 16 (Baijiania).

[Siraitieae+[Momordiceae+[Joliffieae+[Bryonieae+[Sicyoeae+[Schizopeponeae+[Coniandreae+[Benincaseae+Cucurbiteae]]]]]]]]

Siraitieae H. Schaef. et S. S. Renner in Taxon 60: 130. 1 Feb 2011

1/5. Siraitia (5; S. grosvenorii, S. siamensis, S. sikkimensis, S. silomaradjae: eastern Himalayas, southern China, Thailand, Vietnam, Malesia; S. africana: in tropical Africa). – Twining herbs. Tendrils zanonioid, apically bifid. Black to yellow glandular hairs frequent. Stamens five or 2+2+1, adnate at base to corolla tube. Filaments separate. Anthers monothecal. Fruit baccate. n = 14.

[Momordiceae+[Joliffieae+[Bryonieae+[Sicyoeae+[Schizopeponeae+[Coniandreae+[Benincaseae+Cucurbiteae]]]]]]]

Momordiceae H. Schaef. et S. S. Renner in Taxon 60: 130. 1 Feb 2011

1/35–40. Momordica (35–40; tropical and subtropical Africa, the Arabian Peninsula, tropical Asia to eastern Queensland). – Usually lianas or twining herbs (rarely shrubs). Tendrils simple or apically bifid. Stamens three or two, adnate to corolla tube (epipetalous). Filaments separate. Two anthers dithecal and one anther monothecal, or one anther trithecal and one dithecal. Fruit indehiscent or capsular (dehiscing by three valves or irregularly). Aril often present. n = 11, 14.

[Joliffieae+[Bryonieae+[Sicyoeae+[Schizopeponeae+[Coniandreae+[Benincaseae+Cucurbiteae]]]]]]

Joliffieae Schrad. in Linnaea 12: 402. Apr-Sep 1838

3/9. Cogniauxia (2; C. podolaena, C. trilobata; Central Africa), Telfairia (3; T. batesii, T. occidentalis, T. pedata; tropical Africa), Ampelosycios (4; A. humblotii, A. major, A. meridionalis, A. scandens; Madagascar). – Tropical Africa, Madagascar. Lianas or twining herbs. Tendrils simple or bifid. Stamens three (or five). Fruit fleshy. n = 12 (Telfairia). Eleostearic acid (isomer of punicic acid) present.

[Bryonieae+[Sicyoeae+[Schizopeponeae+[Coniandreae+[Benincaseae+Cucurbiteae]]]]]

Bryonieae Dumort., Fl. Belg.: 54. 1827

3/18. Austrobryonia (4; A. argillicola, A. centralis, A. micrantha, A. pilbarensis; northwestern and central Australia), Bryonia (13; Europe, the Canary Islands, the Mediterranean, North Africa, Southwest and Central Asia), Ecballium (1; E. elaterium; the Mediterranean). – Europe, the Mediterranean, North Africa, the Canary Islands, Southwest and Central Asia, northwestern and central Australia. Twining herbs. Tendrils simple or absent. Stamens three. Fruit a berry (in Ecballium elaterium ejecting seeds by elastic contraction). n = 9 (Ecballium), 10 (Bryonia).

[Sicyoeae+[Schizopeponeae+[Coniandreae+[Benincaseae+Cucurbiteae]]]]

Sicyoeae Schrad. in Linnaea 12: 407. Apr-Sep 1838 [‘Secyoideae’]

12/244. Nothoalsomitra (1; N. suberosa; southeastern Queensland); Luffa (5; L. acutangula, L. aegyptiaca, L. cylindrica, L. operculata, L. sepium; tropical regions on both hemispheres); Trichosanthes (83; tropical Asia to Queensland and islands in the Pacific), Hodgsonia (2; H. heteroclita, H. macrocarpa; Assam, Bhutan, southern China, Burma, Thailand, Indochina, Malesiatropical Asia), Linnaeosicyos (1; L. amara; Hispaniola), Echinocystis (1; E. lobata; eastern North America), Marah (8; western and southeastern United States, Mexico), Frantzia (6; F. panamensis, F. pittieri, F. tacaco, F. talamancensis, F. venosa, F. villosa; Central America), Sicyos (67; eastern Australia, Tasmania, Norfolk Island, Lord Howe, New Zealand, islands in the Pacific incl. the Hawaiian Islands, tropical and subtropical America), Hanburia (7; H. caracasana, H. grisebachii, H. mexicana, H. oerstedii, H. parviflora, H. spectabilis, H. subcyclanthera; Mexico, Central America to tropical South America), Cyclanthera (42; southwestern United States, Mexico, tropical America), Echinopepon (21; southern United States to northern Argentina, with their highest diversity in western Mexico). – Pantropical. Lianas or twining herbs. Tendrils simple or bifid to octafid. Multicellular nectariferous hairs present on hypanthium. Stamens two to five. Anther thecae often curved, sigmoid, convolute or fused to a flat or folded ring, dehiscing by continuous slit. Pistil sometimes consisting of one carpel. Endosperm haustorium in Sicyos edulis up to 19 mm long. Fruit dry or fleshy, indehiscent, explosively dehiscent or a pyxidium. n = 11–16(–44).

[Schizopeponeae+[Coniandreae+[Benincaseae+Cucurbiteae]]]

Schizopeponeae C. Jeffrey in Kew Bull. 17: 475. 9 Apr 1964

2/9. Schizopepon (8; northern India, the Himalayas, Tibet, China, eastern Siberia, Japan, Burma), Herpetospermum (1; H. pedunculosum; eastern Himalayas to Burma and southwestern China). – The Himalayas, Tibet, China, Japan. Twining herbs. Tendrils bifid or trifid. Stamens three. Pollen grains sometimes triporate. Fruit indehiscent or capsular. n = 10 (Schizopepon), 11 (Herpetospermum).

[Coniandreae+[Benincaseae+Cucurbiteae]]

Coniandreae Endl. ex M. Roem., Fam. Nat. Syn. Monogr. 2: 6. Dec 1846

19/175–195. Bambekea (1; B. racemosa; western and central tropical Africa), Eureiandra (8; tropical and subtropical Africa, Madagascar, Socotra), Dendrosicyos (1; D. socotranus; Yemen, Socotra), Seyrigia (5; S. bosseri, S. gracilis, S. humbertii, S. marnieri, S. multiflora; southern and southwestern Madagascar), Trochomeriopsis (1; T. diversifolia; Madagascar), Halosicyos (1; H. ragonesei; central Argentina), Cucurbitella (1; C. asperata; Brazil, Paraguay, Uruguay, Bolivia, Argentina), Corallocarpus (13–16; tropical Africa, Madagascar, the Arabian Peninsula, Pakistan, India), Kedrostis (c 20; tropical and subtropical Africa, Madagascar, the Arabian Peninsula, India, Sri Lanka, West Malesia), Ceratosanthes (12; Central America to northern Argentina), Doyerea (1; D. emetocathartica; tropical America), Gurania (40–45; tropical America), Psiguria (12–17; Central America, tropical South America; incl. Helmontia?), Helmontia (4; H. cardiophylla, H. leptantha, H. simplicifolia, H. trujilloi; Venezuela, Guyana, Brazil; in Psiguria?), Melothrianthus (1; M. smilacifolius; Brazil), Wilbrandia (c 15; Brazil, Paraguay, Argentina), Apodanthera (30–35; tropical and subtropical America), Tumamoca (2; T. macdougalii: near Tucson in Arizona, Sonora in Mexico; T. mucronata: Zacatecas in central Mexico), Ibervillea (10–11; Texas to Guatemala). – Tropical and subtropical Africa, Madagascar, Socotra, the Arabian Peninsula, India, tropical Asia, southern United States to northern Argentina. Usually lianas or twining herbs (rarely trees). Tendrils simple or bifid or trifid (rarely absent). Stamens two, three or five. Thecae often curved, convolute etc. Pollen grains sometimes shed in tetrads. Fruit baccate. Testal hypodermis not sclerotic. n = 12 or 13 (Kedrostis), 13 (Seyrigia, Corallocarpus), 14 (Apodanthera).

[Benincaseae+Cucurbiteae]

Benincaseae Ser. in Mém. Soc. Phys. Genève 3(1): 25. 1825 [‘Beninsaceae’]

24/230–235. Citrullus (4; C. colocynthis, C. ecirrhosus, C. lanatus, C. rehmii; eastern Mediterranean, North Africa, tropical and southern Africa, western Asia), Peponium (c 20; tropical and subtropical Africa, Madagascar, the Seychelles), Lagenaria (6; L. abyssinica, L. breviflora, L. guineensis, L. rufa, L. siceraria, L. sphaerica; tropical Africa, Madagascar, one species, L. siceraria, pantropical), Acanthosicyos (1; A. horridus; Angola, Namibia, Botswana, South Africa), Raphidiocystis (4; R. brachypoda, R. chrysocoma, R. jeffreyana, R. phyllocalyx; tropical Africa, Madagascar), Cephalopentandra (1; C. ecirrhosa; northeastern tropical Africa), Lemurosicyos (1; L. variegata; Madagascar), Solena (3; S. amplexicaulis, S. delavayi, S. heterophylla; tropical Asia), Borneosicyos (1; B. simplex; Sarawak, Sabah), Benincasa (2; B. hispida; New Caledonia, New Ireland, New Guinea, Queensland; B. fistulosa: known only from cultivation), Ctenolepis (1; C. lucorum; Madagascar), Dactyliandra (2; D. welwitschii: the Namib desert in Angola and Namibia, the Thar desert in Pakistan and India; D. nigrescens: Kenya), Khmeriosicyos (1; K. harmandii; Cambodia; probably extinct), Papuasicyos (c 8; New Guinea), Scopellaria (2; S. diversifolia, S. marginata; Yunnan, Southeast Asia, West Malesia to the Philippines), Trochomeria (8; subtropical and tropical Africa), Indomelothria (2; I. blumei, I. chlorocarpa; Southeast Asia, West Malesia), Melothria (16; southeastern United States, Mexico, Central America, tropical South America; one species, M. mannii, in western tropical Africa and in Central America and tropical South America), Ruthalicia (2; R. eglandulosa, R. longipes; tropical West Africa), Muellerargia (2; M. jeffreyana: Madagascar; M. timorensis: the Lesser Sunda Islands, Timor, tropical northern Australia), Cucumis (c 55; subtropical and tropical regions in the Old World), Zehneria (c 60; tropical regions in the Old World), Diplocyclos (4–5; D. decipiens, D. leiocarpus, D. palmatus, D. schliebenii, D. tenuis; subtropical and tropical Africa, tropical Asia, tropical Australia), Coccinia (c 25; tropical and southern Africa, one species, C. cordifolia, also in tropical Asia). – Eastern Mediterranean, Africa, Madagascar, the Seychelles, Aldabra, western and tropical Asia to New Guinea, Australia and Melanesia, islands in the Pacific, southeastern United States to South America, with their highest diversity in tropical Africa and Madagascar. Usually lianas or twining herbs (rarely shrubs). Tendrils simple, 2–5-fid or absent. Paired spines present at nodes in Acanthosicyos. Stamens usually three (sometimes two, four or five). Two anthers dithecal and one monothecal or all dithecal or all monothecal. Thecae often curved or convolute, etc. Fruit usually baccate. n = 7, 11, 12, 24. Mitochondrial coxI intron present.

Cucurbiteae Dumort., Fl. Belg.: 54. 1827

11/95–110. Polyclathra (1 or 6; P. cucumerina; Mexico, Central America), Peponopsis (1; P. adhaerens; Mexico), Cucurbita (17; tropical and subtropical America), Calycophysum (5; C. gracile, C. pedunculatum, C. spectabile, C. villosum, C. weberbaueri; northwestern tropical South America), Sicana (3; S. odorifera, S. sphaerica, S. trinitensis; Central America, the West Indies), Penelopeia (2; P. sphaerica, P. suburceolata; Hispaniola), Tecunumania (1; T. quetzalteca; Mexico, Guatemala, Costa Rica), Schizocarpum (11; Mexico, Guatemala), Cionosicyos (4; C. excisus, C. guabubu, C. macranthus, C. pomiformis; Central America, Cuba, Jamaica), Abobra (1; A. tenuifolia; southern Brazil, Uruguay, Argentina), Cayaponia (450–60; southern United States, Mexico, Central America, tropical South America, one species, C. noronhae, endemic on Fernando de Noronha, one species, C. africana, in western and central tropical Africa and Madagascar). – Southern United States, Mexico, the Caribbean, Central and South America, tropical Africa, Madagascar. Lianas or twining herbs. Tendrils simple or 2–7-fid. Extrafascicular sieve tubes in Cucurbita present inside cylinder of “pericyclic” fibres. Stamens usually three (sometimes two or four). Thecae often reflexed, convolute, etc. Pollen grains triporate to periporate (sometimes up to 200 μm). Fruit small dry and indehiscent, or medium-sized or large pepos, or dry and capsular (dehiscing with several valves). n = 20 (Cucurbita, Sicana).

Phylogeny (simplified) of Cucurbitaceae based on DNA data (Schaeffer & Renner 2011).

Datiscales Bercht. et J. Presl, Přir. Rostlin: 217. Jan-Apr 1820 [‘Datisceae’]; Datiscineae Engl., Syllabus, ed. 2: 156. Mai 1898

Distribution Crete to Turkey and the Caucasus, western Himalayas, northern California to northwestern Mexico.

Habit Polygamomonoecious, dioecious or androdioecious, perennial herbs. Superficially similar in habit to Cannabis (Cannabaceae).

Vegetative anatomy Root nodules containing nitrogen fixing endosymbiotic actinobacteria (Frankia). Phellogen ab initio superficial. Medullary vascular bundles present. Cambium and wood elements not storied. Vessel elements with simple perforation plates; lateral pits alternate, simple pits. Imperforate tracheary xylem elements libriform fibres with simple pits, non-septate? Wood rays absent? Axial parenchyma paratracheal? Sieve tube plastids S type. Nodes 1:3, unilacunar with three leaf traces. Tanniniferous sacs present. Crystals?

Trichomes Hairs unicellular or multicellular, uniseriate, simple; glandular hairs?

Leaves Alternate (spiral), deeply pinnately lobed to pinnately compound (imparipinnate), with conduplicate ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection? Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids? Domatia as pockets. Mesophyll without sclerenchymatous idioblasts. Leaf margin serrate.

Flowers Actinomorphic. Epigyny. Sepals three to nine (to ten), with valvate aestivation (in male flowers), persistent, free. Petals absent. Nectary absent. Disc absent.

Androecium Stamens in bisexual flowers three to five, in male flowers eight to c. 25, outer stamens often antesepalous. Filaments very short, free from each other and from tepals. Anthers straight, basifixed, non-versatile, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia usually absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate, shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, rugulate.

Gynoecium Pistil composed of three to five (to eight) antesepalous carpels connate in lower part; carpels strongly developed, ventrally forming roof-like structure above locule, this forming perianth tube base; calyx tube with widely separated stylodia inserted on roof edge. Ovary inferior, unilocular. Stylodia three to five (to eight), subulate, deeply bifid; compitum possibly present. Stigmas papillate, Dry type. Pistillodium absent.

Ovules Placentation parietal. Ovules c. 30 to more than 100 per carpel, anatropous, pendulous to horizontal, bitegmic, crassinucellar. Micropyle bistomal. Outer integument two cell layers thick. Inner integument two cell layers thick. Parietal tissue three to five cell layers thick. Nucellar cap two or three cell layers thick. Megagametophyte disporous, 8-nucleate, Allium type. Antipodal cells persistent. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad.

Fruit A membranous septicidal? capsule, apically dehiscing between persistent stylodia.

Seeds Aril absent. Funicle surrounded by collar formed as extension of testa. Operculum formed by micropyle and hilum, without surrounding annulus of collar cells. Seed coat exotestal. Exotestal cells with honeycomb-like arrangement, with strongly thickened and lignified inner walls. Endotesta absent. Exotegmic cells large, cuboid. Endotegmen absent. Perisperm not developed. Endosperm sparse with oil and aleurone, or absent. Embryo small, straight, well differentiated, chlorophyll? Cotyledons two, oily. Germination phanerocotylar.

Phytochemistry B-ring-nonsubstituted and 2’-hydroxylated flavonols (kaempferol, quercetin) and cucurbitacins present. Glucosides of unusual flavonols, galangin (5, 7-dihydroxyflavonol), 7-O-methylgalangin, datiscetin (3, 5, 7, 2’-tetrahydroxyflavonol) and 7-O-methyldatiscetin present. Ellagic acid, tannins, proanthocyanidins, and cyanogenic compounds not found.

Fossils Fossil wood, Tetrameleoxylon prenudiflora, has been described from the Maastrichtian Deccan Intertrappean Beds in India.

Vegetative anatomy Phellogen ab initio superficial. Vessel elements with simple perforation plates; lateral pits alternate, simple pits. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, non-septate. Wood rays multiseriate, heterocellular. Axial parenchyma paratracheal scanty, aliform, confluent, vasicentric, or banded. Wood soft, sometimes fluorescent. Wood elements often storied. Sieve tube plastids S type. Nodes usually 3:1–2, trilacunar with one or two leaf traces from each lateral leaf gap. Branched sclereids and stone cells present in cortex and medulla in Octomeles. Crystals?

Leaves Alternate (spiral), simple, entire, asymmetrical, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection? Venation palmate. Stomata anomocytic. Cuticular wax crystalloids absent? Mesophyll in at least Octomeles with sclerenchymatous H-shaped idioblasts (reaching from one epidermis to next). Leaf margin usually serrate (rarely entire).

Flowers Actinomorphic. Hypanthium present. Epigyny. Sepals four or six to eight, with valvate aestivation, free or connate (postgenitally coherent). Petals usually absent (in male flowers of Octomeles six to eight, small, with valvate aestivation, free, inserted on sepals). Nectaries possibly present inside male floral tube in Octomeles. Disc absent or present above ovary.

Androecium Stamens four or six to eight, obhaplostemonous, alternisepalous, antepetalous. Filaments flattened to narrowly elongate, free from each other and from tepals, inflexed in bud. Anthers straight or curved, basifixed, non-versatile, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Female flowers sometimes with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Pistil composed of (three or) four or six to eight carpels connate in lower part; carpels strongly developed ventrally forming roof-like structure above locule, this forming perianth tube base; calyx tube with widely separated stylodia inserted on roof edge. Compitum possibly present in Octomeles. Ovary inferior, unilocular, septate. Stylodia (three or) four or six to eight, short, straight, free. Stigmas entire, decurrent to clavate or capitate, type? Male flowers sometimes with pistillodium.

Ovules Placentation parietal, protruding-diffuse; placentae in Octomeles bilobate. Ovules c. 20 to more than 100 per carpel, anatropous, pendulous or horizontal, bitegmic, crassinucellar. Micropyle bistomal. Outer integument approx. two cell layers thick. Inner integument approx. two cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A septicidal capsule, in Tetrameles apically dehiscing between persistent stylodia and along sides, in Octomeles separating into two layers outer of which being detached.

Seeds Aril absent? Funicle surrounded by collar formed as extension of testa. Operculum formed by micropyle and hilum, without surrounding annulus of cells. Seed coat exotestal. Exotestal cells with honeycomb-like arrangement, with strongly thickened and lignified inner walls? Endotesta? Tegmen? Perisperm not developed. Endosperm very sparse or absent. Embryo straight, well differentiated, chlorophyll? Cotyledons two. Germination?

Phytochemistry Insufficiently known. Flavonols (kaempferol, quercetin) and cucurbitacins present. Tannins and cyanogenic compounds not found.

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”The Nitrogen Fixing clade”

CUCURBITALES Juss. ex Bercht. et J. Presl

ANISOPHYLLEACEAE Ridl.

APODANTHACEAE (R. Br.) Tiegh. ex Takht.

BEGONIACEAE C. Agardh

CORIARIACEAE DC.

CORYNOCARPACEAE Engl.

CUCURBITACEAE Juss.

DATISCACEAE Bercht. et J. Presl

TETRAMELACEAE (Warb.) Airy Shaw